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<document ID-DOI="http://dx.doi.org/10.3897/vz.72.e89647" ID-GBIF-Dataset="e83eba07-b611-4b55-b94a-96f2059d0efd" ID-Pensoft-Pub="2625-8498-72-997" ID-Pensoft-UUID="F478C7D1544650D99649E511AB6CED15" ID-ZooBank="0CA196A200E04E0AB21611AD5B3856EB" ModsDocID="2625-8498-72-997" checkinTime="1668159806196" checkinUser="pensoft" docAuthor="Kraus, Fred, Kaiser, Hinrich &amp; O'Shea, Mark" docDate="2022" docId="2012093A92CE51D2A5E707D6D6CF5924" docLanguage="en" docName="VertZool 72: 997-1034" docOrigin="Vertebrate Zoology 72" docPubDate="2022-11-10" docSource="http://dx.doi.org/10.3897/vz.72.e89647" docTitle="Toxicocalamus loriae" docType="treatment" docVersion="2" id="F478C7D1544650D99649E511AB6CED15" lastPageNumber="997" masterDocId="F478C7D1544650D99649E511AB6CED15" masterDocTitle="Hidden diversity in semi-fossorial Melanesian forest snakes: A revision of the Toxicocalamus loriae complex (Squamata, Elapidae) from New Guinea" masterLastPageNumber="1034" masterPageNumber="997" pageNumber="997" updateTime="1668247209537" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Hidden diversity in semi-fossorial Melanesian forest snakes: A revision of the Toxicocalamus loriae complex (Squamata, Elapidae) from New Guinea</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kraus, Fred</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0003-4194-4959</mods:nameIdentifier>
<mods:affiliation>Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kaiser, Hinrich</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-0001-9428</mods:nameIdentifier>
<mods:affiliation>Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA &amp; Department of Vertebrate Zoology, Leibniz-Institut zur Analyse des Biodiversitaetswandels, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany</mods:affiliation>
<mods:nameIdentifier type="email">chalcopis@yahoo.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>O'Shea, Mark</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-1566-7460</mods:nameIdentifier>
<mods:affiliation>Faculty of Science and Engineering, University of Wolverhampton, Wulfruna Street, Wolverhampton, West Midlands WV 1 1 LY, United Kingdom</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>Vertebrate Zoology</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-11-10</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>72</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>997</mods:start>
<mods:end>1034</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/vz.72.e89647</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/vz.72.e89647</mods:identifier>
<mods:identifier type="Pensoft-Pub">2625-8498-72-997</mods:identifier>
<mods:identifier type="ZooBank">0CA196A200E04E0AB21611AD5B3856EB</mods:identifier>
<mods:identifier type="Pensoft-UUID">F478C7D1544650D99649E511AB6CED15</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="204928494" LSID="urn:lsid:plazi:treatment:2012093A92CE51D2A5E707D6D6CF5924" httpUri="http://treatment.plazi.org/id/2012093A92CE51D2A5E707D6D6CF5924" lastPageNumber="997" pageId="0" pageNumber="997">
<subSubSection pageId="0" pageNumber="997" type="nomenclature">
<paragraph pageId="0" pageNumber="997">
<taxonomicName LSID="2012093A-92CE-51D2-A5E7-07D6D6CF5924" authority="(Boulenger, 1898)" baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">Toxicocalamus loriae (Boulenger, 1898)</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="description">
<paragraph pageId="0" pageNumber="997">
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal views of heads of Toxicocalamus loriae Group snakes, presented as both photographic and line-drawing illustrations. A, A ' holotype of T. loriae (MSNG 29141), Haveri, Bartholomew Range, Central Province, PNG. B, B ' lectotype of T. nymani comb. nov. (UUMZ 290 / 2387), Sattelberg, Huon Peninsula, Morobe Province, PNG. C, C ' lectotype of T. loennbergii comb. nov. (BMNH 1946.1.18.24), Fakfak, Onin Peninsula, West Papua Province, Indonesia. D, D ' holotype of T. lamingtoni comb. nov. (AMS R 9351), Mount Lamington, Oro Province, PNG. Images not to scale. Scale abbreviations: anterior temporals (AT), frontal (F), internasals (IN), nasals (N), parietals (P), prefrontals (PF), postoculars (PO), preoculars (PR), posterior temporals (PT), rostral (R), supraoculars (SO)." figureDoi="10.3897/vz.72.e89647.figure1" httpUri="https://binary.pensoft.net/fig/768167" pageId="0" pageNumber="997">
Figs 1A, A ', 2A,
<normalizedToken originalValue="A’">A'</normalizedToken>
, B,
<normalizedToken originalValue="B’">B'</normalizedToken>
, 3A,
<normalizedToken originalValue="A’">A'</normalizedToken>
, 4A, B, 5
</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="reference_group">
<paragraph pageId="0" pageNumber="997">
<taxonomicName authorityName="Kraus &amp; Kaiser &amp; O’Shea" authorityYear="2022" baseAuthorityName="Kraus &amp; Kaiser &amp; O’Shea" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Apistocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apistocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">Apistocalamus loriae</taxonomicName>
Boulenger, 1898: 705.
</paragraph>
<paragraph pageId="0" pageNumber="997">
<taxonomicName authorityName="Kraus &amp; Kaiser &amp; O’Shea" authorityYear="2022" baseAuthorityName="Kraus &amp; Kaiser &amp; O’Shea" baseAuthorityYear="2022" class="Reptilia" family="Elapidae" genus="Apistocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apistocalamus Pratti" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="Pratti">Apistocalamus Pratti</taxonomicName>
Boulenger, 1904: 451.
</paragraph>
<paragraph pageId="0" pageNumber="997">
<taxonomicName authorityName="Boulenger" authorityYear="1908" class="Reptilia" family="Elapidae" genus="Apisthocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apisthocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">Apisthocalamus loriae</taxonomicName>
- Boulenger, 1908: 249.
</paragraph>
<paragraph pageId="0" pageNumber="997">
<taxonomicName authorityName="Boulenger" authorityYear="1908" class="Reptilia" family="Elapidae" genus="Apisthocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apisthocalamus pratti" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="pratti">Apisthocalamus pratti</taxonomicName>
- Boulenger, 1908: 249.
</paragraph>
<paragraph pageId="0" pageNumber="997">
<taxonomicName class="Reptilia" family="Elapidae" genus="Apisthocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apisthocalamus prattii" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="prattii">Apisthocalamus prattii</taxonomicName>
- Barbour, 1912: 201.
</paragraph>
<paragraph pageId="0" pageNumber="997">
<taxonomicName authorityName="Kraus &amp; Kaiser &amp; O’Shea" authorityYear="2022" baseAuthorityName="Kraus &amp; Kaiser &amp; O’Shea" baseAuthorityYear="2022" class="Reptilia" family="Elapidae" genus="Apistocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apistocalamus pratti" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="pratti">Apistocalamus pratti</taxonomicName>
- McDowell, 1967: 537.
</paragraph>
<paragraph pageId="0" pageNumber="997">
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus (Apistocalamus) loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae" subGenus="Apistocalamus">Toxicocalamus (Apistocalamus) loriae</taxonomicName>
- McDowell, 1969: 455.
</paragraph>
<paragraph pageId="0" pageNumber="997">
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">Toxicocalamus loriae</taxonomicName>
Clade 1 -
<bibRefCitation DOI="https://doi.org/10.1111/zoj.12423" author="Strickland, JL" journalOrPublisher="Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin" pageId="0" pageNumber="997" refId="B44" refString="Strickland, JL, Carter, S, Kraus, F, Parkinson, CL, 2016. Snake evolution in Melanesia: origin of the Hydrophiinae (Serpentes, Elapidae) and the evolutionary history of the enigmatic New Guinean elapid Toxicocalamus. Zoological Journal of the Linnean Society 178: 663--678. https://doi.org/10.1111/zoj.12423" title="Snake evolution in Melanesia: origin of the Hydrophiinae (Serpentes, Elapidae) and the evolutionary history of the enigmatic New Guinean elapid Toxicocalamus. Zoological Journal of the Linnean Society 178: 663 -- 678." url="https://doi.org/10.1111/zoj.12423" year="2016">Strickland et al., 2016</bibRefCitation>
: 671.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="holotype and collection">
<paragraph pageId="0" pageNumber="997">Holotype and collection.</paragraph>
<paragraph pageId="0" pageNumber="997">
MSNG 29141, an adult male from Haveri, British New Guinea (vicinity of
<geoCoordinate degrees="9.40" direction="south" orientation="latitude" precision="555" value="-9.4">9.40°S</geoCoordinate>
,
<geoCoordinate degrees="147.60" direction="east" orientation="longitude" precision="555" value="147.6">147.60°E</geoCoordinate>
). The specimen was collected by the Italian anthropologist, ethnographer, explorer, and naturalist Lamberto Loria (1855-1913), who travelled in British New Guinea from 1889-1897 and sent a series of zoological specimens to the Museo Civico di Storia Naturale in Genoa, Italy. Based on
<normalizedToken originalValue="Loria’s">Loria's</normalizedToken>
diary accounts of his travels (see
<bibRefCitation author="Dimpflmeier, F" journalOrPublisher="Lares" pageId="0" pageNumber="997" pagination="103 - 124" refId="B13" refString="Dimpflmeier, F, 2014. Itinerari e tappe di Lamberto Loria nella Nuova Guinea Britannica. Lares 80: 103 - 124" title="Itinerari e tappe di Lamberto Loria nella Nuova Guinea Britannica." volume="80" year="2014">Dimpflmeier 2014</bibRefCitation>
,
<bibRefCitation author="Dimpflmeier, F" journalOrPublisher="Lares" pageId="0" pageNumber="997" publicationUrl="https://www.berose.fr/article1755.html" refId="B14" refString="Dimpflmeier, F, 2019. From Italy to British New Guinea and back: The life and (field)work of Lamberto Loria. In: Berose - Encyclopedie Internationale des Histoires de l'Anthropologie, Paris, France. Available at: https://www.berose.fr/article1755.html" title="From Italy to British New Guinea and back: The life and (field) work of Lamberto Loria. In: Berose - Encyclopedie Internationale des Histoires de l'Anthropologie, Paris, France. Available at:" url="https://www.berose.fr/article1755.html" year="2019">2019</bibRefCitation>
), the specimen was most likely collected during an excursion on 28 June 1893 to the southern versant of the Owen Stanley Range a point-to-point distance of 50 km east of Port Moresby. Thus,
<bibRefCitation author="Boulenger, GA" journalOrPublisher="Evolution" pageId="0" pageNumber="997" refId="B6" refString="Boulenger, GA, 1898. An account of the reptiles and batrachians collected by Dr. L. Loria in British New Guinea. Annali del Museo Civico di Storia Naturale de Genova, Seria 2 18: 694-710." title="An account of the reptiles and batrachians collected by Dr. L. Loria in British New Guinea. Annali del Museo Civico di Storia Naturale de Genova, Seria 2 18: 694 - 710." year="1898">
<normalizedToken originalValue="Boulenger’s">Boulenger's</normalizedToken>
(1898)
</bibRefCitation>
estimate of 1889-1892 for the collection he described appears to have been at least a year short. The species was described in Series 2, Volume 18 of the Annali del Museo Civico di Storia Naturale di Genova, which is an 1897 volume that was published in March 1898 (
<bibRefCitation author="Poggi, R" journalOrPublisher="Annali del Museo Civico di Storia Naturale &quot; Giacomo Doria &quot;" pageId="0" pageNumber="997" pagination="1 - 629" refId="B38" refString="Poggi, R, 2010. Gli &quot;Annali&quot; pubblicati dal Museo Civico di Storia Naturale &quot;Giacomo Doria&quot; di Genova, storia del periodico ed indice generali dei primo cento volumi (1870-2009). Annali del Museo Civico di Storia Naturale &quot;Giacomo Doria&quot; 101: 1 - 629" title="Gli &quot; Annali &quot; pubblicati dal Museo Civico di Storia Naturale &quot; Giacomo Doria &quot; di Genova, storia del periodico ed indice generali dei primo cento volumi (1870 - 2009)." volume="101" year="2010">Poggi 2010</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="etymology">
<paragraph pageId="0" pageNumber="997">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="997">
Named by
<bibRefCitation author="Boulenger, GA" journalOrPublisher="Evolution" pageId="0" pageNumber="997" refId="B6" refString="Boulenger, GA, 1898. An account of the reptiles and batrachians collected by Dr. L. Loria in British New Guinea. Annali del Museo Civico di Storia Naturale de Genova, Seria 2 18: 694-710." title="An account of the reptiles and batrachians collected by Dr. L. Loria in British New Guinea. Annali del Museo Civico di Storia Naturale de Genova, Seria 2 18: 694 - 710." year="1898">Boulenger (1898)</bibRefCitation>
for the collector of the holotype. The description was published in English.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="diagnosis">
<paragraph pageId="0" pageNumber="997">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="997">
A modestly sized member of the
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
Group (maximum SVL in males 490 mm, in females 440 mm) with the following unique combination of characters: two scales covering vent; three infralabials contacting anterior genial; posterior genials separated by two intergenials, one in front of the other; intergenials widest anteriorly. Elongate preocular, at least twice as long as wide, contacting nasal but not internasal; one (in 58% of specimens) or two (42%) postoculars; two (55%) or three (45%) posterior temporals; ventral scale count not sexually dimorphic, 162-197 ventrals in 15 males, 172-190 in four females; subcaudal scale count sexually dimorphic without overlap, 40-50 in males, 28-33 in females; SCR 17.1-21.5% in males, 12.5-16.1% in females; pale markings on prefrontals absent (67%), small or vaguely developed (28%), or present (5%), not obviously correlated with body size; tail spine white, paler than the rest of the tail; venter uniformly yellow, yellow with a mid-ventral row of brown spots on each ventral, with a few brown spots scattered down the venter, or with each spot expanded into a brown bar across the anterior of each ventral.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="comparisons">
<paragraph pageId="0" pageNumber="997">Comparisons with other species.</paragraph>
<paragraph pageId="0" pageNumber="997">
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
is distinguished from all other members of the
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
Group in having the posterior genials separated by two intergenials that are aligned in the midline, one in front of the other, with each scale widest anteriorly. All other species are characterized by having the posterior genials separated by only a single intergenial that is widest posteriorly (typically) or centrally (occasionally). As noted above, in the other species there will rarely be two (or three in AMS R23069) intergenials, but the anterior one is either tiny or the two scales occur side-by-side at the posterior portion of the posterior genials; the rarity of these anomalous features suggests they are developmental oddities.
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
may be further distinguished from
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nigrescens" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nigrescens">
<emphasis italics="true" pageId="0" pageNumber="997">T. nigrescens</emphasis>
</taxonomicName>
by its smaller body size (maximum SVL in males 490 mm vs. 635 mm in
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nigrescens" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nigrescens">
<emphasis italics="true" pageId="0" pageNumber="997">T. nigrescens</emphasis>
</taxonomicName>
), and yellow ventral colouration (grey in
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nigrescens" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nigrescens">
<emphasis italics="true" pageId="0" pageNumber="997">T. nigrescens</emphasis>
</taxonomicName>
). It can be distinguished from
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
</taxonomicName>
by preoculars that are in contact with the nasals (separated by prefrontal contact with second supralabial in
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
</taxonomicName>
) and its venter that is uniformly yellow or yellow with a mid-ventral row of brown spots or short bars (venter grey or yellow with each ventral scale barred anteriorly with darker grey in
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="redescription">
<paragraph pageId="0" pageNumber="997">Redescription of the holotype.</paragraph>
<paragraph pageId="0" pageNumber="997">
An adult male, with tongue extending from right side of mouth; 490 mm SVL, tail tip missing but TL&gt; 90 mm. Rostral, internasals, and anterior portions of nasals crossed by series of linear impressions, as if the snout had been pressed against a screen during preservation. Rostral wider than tall, notched ventromedially; internasals angulate, semi-triangular, wider than long. Prefrontals distinct from preoculars, approximately square but angled laterally and posteriorly, slightly longer than wide (Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal views of heads of Toxicocalamus loriae Group snakes, presented as both photographic and line-drawing illustrations. A, A ' holotype of T. loriae (MSNG 29141), Haveri, Bartholomew Range, Central Province, PNG. B, B ' lectotype of T. nymani comb. nov. (UUMZ 290 / 2387), Sattelberg, Huon Peninsula, Morobe Province, PNG. C, C ' lectotype of T. loennbergii comb. nov. (BMNH 1946.1.18.24), Fakfak, Onin Peninsula, West Papua Province, Indonesia. D, D ' holotype of T. lamingtoni comb. nov. (AMS R 9351), Mount Lamington, Oro Province, PNG. Images not to scale. Scale abbreviations: anterior temporals (AT), frontal (F), internasals (IN), nasals (N), parietals (P), prefrontals (PF), postoculars (PO), preoculars (PR), posterior temporals (PT), rostral (R), supraoculars (SO)." figureDoi="10.3897/vz.72.e89647.figure1" httpUri="https://binary.pensoft.net/fig/768167" pageId="0" pageNumber="997">1A, A</figureCitation>
'), bordered below by preocular and nasal; preoculars elongate, narrower anteriorly, approximately twice as long as tall, bordered anteriorly by nasal, below by second and third supralabials (Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Right and left lateral views of heads of Toxicocalamus loriae Group snakes, presented as both photographic and line-drawing illustrations. A, A ', B, B ' holotype of T. loriae (MSNG 29141). C, C ', D, D ' lectotype of T. nymani comb. nov. (UUMZ 290 / 2387). E, E ', F, F ' lectotype of T. loennbergii comb. nov. (BMNH 1946.1.18.24). and G, G ', H, H ' holotype of T. lamingtoni comb. nov. (AMS R 9351). Images not to scale. Scale abbreviations: anterior genial (AG), anterior temporal (AT), frontal (F), numbered infralabials (IN 1 - 6), internasal (IN), mental (M), nasal (N), parietal (P), posterior genial (PG), postocular (PO), preocular (PR), posterior temporals (PT), numbered supralabials (SL 1 - 6), supraocular (SO)." figureDoi="10.3897/vz.72.e89647.figure2" httpUri="https://binary.pensoft.net/fig/768168" pageId="0" pageNumber="997">2A, A</figureCitation>
', B,
<normalizedToken originalValue="B’">B'</normalizedToken>
). Nasal divided by large naris, with distinct groove above and below centre of naris on right side (region damaged on left side). Postoculars two, irregularly shaped, upper approximately four times larger than lower, combined they occupy approximately same area as eye. Frontal shield-shaped, not fused with supraoculars, its anterior margin may extend slightly forward medially from rest of scale; parietals approximately twice as long as wide. One elongate anterior temporal above fifth and sixth supralabials, separating latter from parietal; three posterior temporals, anteriormost smallest, followed posteriorly by a larger upper temporal and below it a smaller lower temporal, ventralmost temporal abutting posterodorsal margin of sixth supralabial. Supralabials six, third and fourth contacting eye; infralabials six, first three in contact with anterior genial. Mental small, shallow, triangular, wider than long, bordered posteriorly by first supralabials; anterior genials wider than posterior genials, in broad medial contact; posterior genials completely separated by two diamond-shaped intergenials, anterior of which is widest anteriorly, posterior of which is widest centrally; four gulars separate intergenials from first ventral in the midline; an elongate sublabial of a length equal to anterior genial separates posterior genial from fifth infralabial (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Ventral views of heads of Toxicocalamus loriae Group snakes, presented as both photographic and line-drawing illustrations. A, A ' holotype of T. loriae (MSNG 29141). B, B ' lectotype of T. nymani comb. nov. (UUMZ 290 / 2387). C, C ' lectotype of T. loennbergii comb. nov. (BMNH 1946.1.18.24). D, D ' holotype of T. lamingtoni comb. nov. (AMS R 9351). Images not to scale. Scale abbreviations: anterior genials (AG), intergenials (IG), numbered infralabials (IL 1 - 6), mental (M), posterior genials (PG), sublabials (Sb)." figureDoi="10.3897/vz.72.e89647.figure3" httpUri="https://binary.pensoft.net/fig/768169" pageId="0" pageNumber="997">3A, A</figureCitation>
'). Eye relatively small; pupil round.
</paragraph>
<caption doi="10.3897/vz.72.e89647.figure1" httpUri="https://binary.pensoft.net/fig/768167" pageId="0" pageNumber="997" start="Figure 1" startId="F1">
<paragraph pageId="0" pageNumber="997">
<emphasis bold="true" pageId="0" pageNumber="997">Figure 1.</emphasis>
Dorsal views of heads of
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
Group snakes, presented as both photographic and line-drawing illustrations.
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
' holotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
(MSNG 29141), Haveri, Bartholomew Range, Central Province, PNG.
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
' lectotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(UUMZ 290/2387), Sattelberg, Huon Peninsula, Morobe Province, PNG.
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
' lectotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loennbergii" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loennbergii">
<emphasis italics="true" pageId="0" pageNumber="997">T. loennbergii</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(BMNH 1946.1.18.24), Fakfak, Onin Peninsula, West Papua Province, Indonesia.
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
' holotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. lamingtoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="lamingtoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. lamingtoni</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(AMS R9351), Mount Lamington, Oro Province, PNG. Images not to scale. Scale abbreviations: anterior temporals (AT), frontal (F), internasals (IN), nasals (N), parietals (P), prefrontals (PF), postoculars (PO), preoculars (PR), posterior temporals (PT), rostral (R), supraoculars (SO).
</paragraph>
</caption>
<caption doi="10.3897/vz.72.e89647.figure2" httpUri="https://binary.pensoft.net/fig/768168" pageId="0" pageNumber="997" start="Figure 2" startId="F2">
<paragraph pageId="0" pageNumber="997">
<emphasis bold="true" pageId="0" pageNumber="997">Figure 2.</emphasis>
Right and left lateral views of heads of
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
Group snakes, presented as both photographic and line-drawing illustrations.
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
',
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
' holotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
(MSNG 29141).
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
',
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
' lectotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(UUMZ 290/2387).
<emphasis bold="true" pageId="0" pageNumber="997">E</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">E</emphasis>
',
<emphasis bold="true" pageId="0" pageNumber="997">F</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">F</emphasis>
' lectotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loennbergii" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loennbergii">
<emphasis italics="true" pageId="0" pageNumber="997">T. loennbergii</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(BMNH 1946.1.18.24). and
<emphasis bold="true" pageId="0" pageNumber="997">G</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">G</emphasis>
',
<emphasis bold="true" pageId="0" pageNumber="997">H</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">H</emphasis>
' holotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. lamingtoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="lamingtoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. lamingtoni</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(AMS R9351). Images not to scale. Scale abbreviations: anterior genial (AG), anterior temporal (AT), frontal (F), numbered infralabials (IN1-6), internasal (IN), mental (M), nasal (N), parietal (P), posterior genial (PG), postocular (PO), preocular (PR), posterior temporals (PT), numbered supralabials (SL1-6), supraocular (SO).
</paragraph>
</caption>
<caption doi="10.3897/vz.72.e89647.figure3" httpUri="https://binary.pensoft.net/fig/768169" pageId="0" pageNumber="997" start="Figure 3" startId="F3">
<paragraph pageId="0" pageNumber="997">
<emphasis bold="true" pageId="0" pageNumber="997">Figure 3.</emphasis>
Ventral views of heads of
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
Group snakes, presented as both photographic and line-drawing illustrations.
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
' holotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
(MSNG 29141).
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
' lectotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(UUMZ 290/2387).
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
' lectotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loennbergii" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loennbergii">
<emphasis italics="true" pageId="0" pageNumber="997">T. loennbergii</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(BMNH 1946.1.18.24).
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
' holotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. lamingtoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="lamingtoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. lamingtoni</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(AMS R9351). Images not to scale. Scale abbreviations: anterior genials (AG), intergenials (IG), numbered infralabials (IL1-6), mental (M), posterior genials (PG), sublabials (Sb).
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="997">Dorsal scales smooth, not notched posteriorly, without apical pits, in 15-15-15 rows; ventrals 189, each approximately four times as wide as long, tail tip incomplete; two scales covering vent; subcaudals 44+, paired.</paragraph>
<paragraph pageId="0" pageNumber="997">In preservative (122 years after collection), dorsum uniformly medium brown; venter yellow with a mid-ventral row of brown spots. Supralabials pale yellow ventrally, suffused with brown on upper portions of Supralabials 2-5 and on posterior portion of Supralabial 6; small amount of yellow colouration also present on the lower parts of the nasals. Chin and throat yellow, with small area of brown suffusion on posterior portions of infralabials, genials, intergenials, and gulars. Iris black.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="variation">
<paragraph pageId="0" pageNumber="997">Variation.</paragraph>
<paragraph pageId="0" pageNumber="997">The anterior margin of the frontal may be almost straight but usually extends slightly forward of the rest of the scale in the midline of the head. The nasals usually have a distinct groove above and below the posterior margin of each naris; a sample from Mt. Obree has either very shallow grooves or lacks them entirely. One (58%) or two (42%) postoculars, both smaller than or occupying approximately the same area as an eye; two (55%) or three (45%) posterior temporals, positioned either as one larger scale above a smaller one, or with the smallest of three scales positioned anterior to the other two, followed posteriorly by a larger upper and a smaller lower temporal; in either configuration, the lowest temporal scale is abutting the posterodorsal margin of the sixth supralabial. Six (97.4%) or seven (2.6%) supralabials, the third and fourth (97.4%) or the third to fifth (2.6%) supralabials contacting eye; five (2.6%) or six (97.4%) infralabials, the first three (22%) or four (78%) in contact with the anterior genial. The anterior genials are wider than the posterior genials and approximately the same length or slightly longer; anterior genials in broad medial contact except in NMW 23783.1 and USNM 195619, where they are separated along most or all their length by an intergenial; posterior genials invariably separated by two diamond-shaped intergenials, whose anterior width is greatest.</paragraph>
<paragraph pageId="0" pageNumber="997">
Dorsal scales invariably in 15-15-15 rows. Ventrals not sexually dimorphic (
<emphasis italics="true" pageId="0" pageNumber="997">t</emphasis>
14 = -0.2688,
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
= 0.3958), 162-197 (184
<normalizedToken originalValue="±">+/-</normalizedToken>
10) in 15 males, 172-190 (179
<normalizedToken originalValue="±">+/-</normalizedToken>
7) in four females; subcaudals sexually dimorphic without overlap, 40-50 in males, 28-33 in females; SCR sexually dimorphic without overlap, 17.1-21.5% (19.4
<normalizedToken originalValue="±">+/-</normalizedToken>
1.4%) in males, 12.5-16.1% (14.6
<normalizedToken originalValue="±">+/-</normalizedToken>
1.0%) in females. Tail tipped by a blunt to pointed conical spine. Maximum SVL 490 mm in males and 440 mm in females, adult TLR = 13.4-19.1% (16.6
<normalizedToken originalValue="±">+/-</normalizedToken>
1.7%) and 10.8-17.5% (14.1
<normalizedToken originalValue="±">+/-</normalizedToken>
3.3%), in males and females, respectively. The current dataset, with only two adult females, indicates that there is no statistically significant male-female difference in SVL (
<emphasis italics="true" pageId="0" pageNumber="997">t</emphasis>
= 0.3034,
<emphasis italics="true" pageId="0" pageNumber="997">p</emphasis>
= 0.3832). TLR in the two juvenile males (PNGM 23158, 24649b) is smaller (12.6% and 13.9%, respectively) than in adults, indicating that in this species, there may be an ontogenetic lengthening of the tail relative to SVL in males. In the four females, this trend appears to be reversed. Even though the sample size is very small, the two subadult females (BPBM 10966, 44892) display TLR values of 12.0% and 11.8%, respectively, with the adult individuals displaying slightly shorter tails relative to their SVLs.
</paragraph>
<paragraph pageId="0" pageNumber="997">
In preservative, the dorsal colouration is uniformly medium brown (84%) or dark brown (11%) except that one specimen (BPBM 10967) is grey-brown mid-dorsally and laterally and has a dorsolateral stripe of medium brown extending the length of the body. The ventral colouration is uniformly pale yellow (37%), yellow with a mid-ventral row of brown spots (32%), yellow with some mid-ventral brown spots that are not present on each scale (11%), or yellow with a brown bar across the anterior portion of each ventral (16%) and with the brown colour more concentrated mid-ventrally into semi-circular spots; one specimen (NMW 27383.2) is uniformly pale brown ventrally, but this may be a discolouration artifact of an originally yellow venter inasmuch as the specimen was preserved in 1904-1906). Lower portions of the supralabials pale yellow, some or all suffused with brown on their upper portions. Yellow markings variably present on nasals, prefrontals, temporals, parietals, and in the nuchal area, virtually absent in the holotype and the sample from Mt. Obree; pale, invariably incomplete nuchal collar, usually limited to lateral markings but in some specimens with small, mid-dorsal yellow spots. Chin and throat pale yellow, usually with a small amount of brown suffusion on portions of the mental, the infralabials, and sometimes on the genials or the anterior intergenial. Conical tail spine with a white (95%) or brown or pale brown (5%) tip. Iris black. Juveniles (
<emphasis italics="true" pageId="0" pageNumber="997">n</emphasis>
= 4) have a brown head with a broken yellow nuchal band and yellow supralabials (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Dorsal and right lateral views of heads of juveniles of Toxicocalamus loriae Group snakes. A, B T. loriae (BPBM 10966), Agaun, Milne Bay Province, PNG. C, D T. nymani comb. nov., spotted form (BPBM 5442), Kalolo, Morobe Province, PNG. E, F T. nymani, dark form (BPBM 23699), Wau, Morobe Province, PNG. G, H T. lamingtoni comb. nov. (AMS R 9352), Mt. Lamington, Oro Province, PNG. I, J T. vertebralis sp. nov. (KU 129086), Wau, Morobe Province, PNG. K, L T. spilorhynchus sp. nov. (AMNH R- 107204), Garaina, Morobe Province, PNG. M, N T. atratus sp. nov. (CAS 118958), Mintima, Chimbu Province, PNG." figureDoi="10.3897/vz.72.e89647.figure4" httpUri="https://binary.pensoft.net/fig/768170" pageId="0" pageNumber="997">4A, B</figureCitation>
). They may or may not have a small yellow spot on each prefrontal.
</paragraph>
<caption doi="10.3897/vz.72.e89647.figure4" httpUri="https://binary.pensoft.net/fig/768170" pageId="0" pageNumber="997" start="Figure 4" startId="F4">
<paragraph pageId="0" pageNumber="997">
<emphasis bold="true" pageId="0" pageNumber="997">Figure 4.</emphasis>
Dorsal and right lateral views of heads of juveniles of
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
Group snakes.
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
(BPBM 10966), Agaun, Milne Bay Province, PNG.
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
, spotted form (BPBM 5442), Kalolo, Morobe Province, PNG.
<emphasis bold="true" pageId="0" pageNumber="997">E</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">F</emphasis>
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
</taxonomicName>
, dark form (BPBM 23699), Wau, Morobe Province, PNG.
<emphasis bold="true" pageId="0" pageNumber="997">G</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">H</emphasis>
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. lamingtoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="lamingtoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. lamingtoni</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(AMS R9352), Mt. Lamington, Oro Province, PNG.
<emphasis bold="true" pageId="0" pageNumber="997">I</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">J</emphasis>
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. vertebralis" order="Squamata" pageId="0" pageNumber="997" rank="species" species="vertebralis">
<emphasis italics="true" pageId="0" pageNumber="997">T. vertebralis</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
(KU 129086), Wau, Morobe Province, PNG.
<emphasis bold="true" pageId="0" pageNumber="997">K</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">L</emphasis>
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. spilorhynchus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="spilorhynchus">
<emphasis italics="true" pageId="0" pageNumber="997">T. spilorhynchus</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
(AMNH R-107204), Garaina, Morobe Province, PNG.
<emphasis bold="true" pageId="0" pageNumber="997">M</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">N</emphasis>
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
(CAS 118958), Mintima, Chimbu Province, PNG.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="997">
In life, field notes described BPBM 19502 as: &quot;Dorsum dark slate-gray, changing to gray-brown laterally. No nuchal collar.&quot; BPBM 19503 was dark olive brown dorsally and slightly lighter dorsolaterally (Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Portraits in life A, B and perimortem ventral views C, D of Toxicocalamus loriae BPBM 19503 A, B, C and BPBM 19502 D from NW side Mt. Obree, Central Province, PNG." figureDoi="10.3897/vz.72.e89647.figure5" httpUri="https://binary.pensoft.net/fig/768171" pageId="0" pageNumber="997">5A, B</figureCitation>
), its venter and supralabials were bright lemon yellow, and its subcaudals were bright lemon yellow with grey margins (Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Portraits in life A, B and perimortem ventral views C, D of Toxicocalamus loriae BPBM 19503 A, B, C and BPBM 19502 D from NW side Mt. Obree, Central Province, PNG." figureDoi="10.3897/vz.72.e89647.figure5" httpUri="https://binary.pensoft.net/fig/768171" pageId="0" pageNumber="997">5C</figureCitation>
). The venter of BPBM 19502 was uniform yellow with a more orange cast than BPBM 19503 and featuring grey mid-ventral spots at the centre of ventrals in the posterior third of body; the subcaudals were lemon yellow, with dark grey margins (Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Portraits in life A, B and perimortem ventral views C, D of Toxicocalamus loriae BPBM 19503 A, B, C and BPBM 19502 D from NW side Mt. Obree, Central Province, PNG." figureDoi="10.3897/vz.72.e89647.figure5" httpUri="https://binary.pensoft.net/fig/768171" pageId="0" pageNumber="997">5D</figureCitation>
).
</paragraph>
<caption doi="10.3897/vz.72.e89647.figure5" httpUri="https://binary.pensoft.net/fig/768171" pageId="0" pageNumber="997" start="Figure 5" startId="F5">
<paragraph pageId="0" pageNumber="997">
<emphasis bold="true" pageId="0" pageNumber="997">Figure 5.</emphasis>
Portraits in life
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
and perimortem ventral views
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
of
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
BPBM 19503
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
and BPBM 19502
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
from NW side Mt. Obree, Central Province, PNG.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="range">
<paragraph pageId="0" pageNumber="997">Range.</paragraph>
<paragraph pageId="0" pageNumber="997">
Known almost exclusively from the southern versant of the Owen Stanley Range of Papua New Guinea, from Tapini, Central Province, in the north to Mt. Dayman, Milne Bay Province, in the south, at elevations from 620-1530 m (Fig.
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Type localities (stars) and other collection sites (circles) of Toxicocalamus loriae Group species. A Map of New Guinea and its satellite islands, showing the positions of the inset maps and the type locality of T. mintoni (magenta), an endemic of Sudest Island, Milne Bay Province, PNG. Scale = 500 km. B Papua New Guinea and adjacent areas of West New Guinea, Indonesia, with white lines demarcating provincial boundaries. Scale = 250 km. C West Papua Province, Indonesia (WP), showing the type locality of T. loennbergii comb. nov. (orange). Other species treated in this report are T. loriae (dark blue), T. lamingtoni comb. nov. (yellow), T. spilorhynchus sp. nov. (red), T. nymani comb. nov. (neon green), T. vertebralis sp. nov. (pink), and T. atratus sp. nov. (light blue). Additional related species in the T. loriae clade are T. goodenoughensis (brown), T. mattisoni (violet), T. nigrescens (black), T. pachysomus (dark green), and T. &quot; Toxicocalamus loriae &quot; Clade 4 (white). The type locality of the recently described dubious taxon T. longhagen (tan) is also indicated. Scale = 250 km. Provinces of PNG are Central (Ce), Chimbu (Ch; also spelled Simbu), Gulf (Gu), Hela (He; created in 2012), Eastern Highlands (EH), East Sepik (ES), Enga (En), Jiwaka (Ji; created in 2012), Madang (Ma), Milne Bay (MB), Morobe (Mo), National Capital District (NCD), Oro (Or; also known as Northern Province); Sandaun (Sa; formerly West Sepik), Southern Highlands (SH), Western (We), and Western Highlands (WH). Provinces of West New Guinea are Highland Papua (HP; created in 2022), Papua (Pa), and South Papua (SP; created in 2022). Sites of sympatry are Wau (W; neon green + pink) with two species and Garaina (G; neon green + red + yellow) with three species." figureDoi="10.3897/vz.72.e89647.figure6" httpUri="https://binary.pensoft.net/fig/768172" pageId="0" pageNumber="997">6B</figureCitation>
). A single specimen (BPBM 44892) is known from Isurava, along the Kokoda Track, on the north side of the Owen Stanley Mountains in Oro Province.
</paragraph>
<caption doi="10.3897/vz.72.e89647.figure6" httpUri="https://binary.pensoft.net/fig/768172" pageId="0" pageNumber="997" start="Figure 6" startId="F6">
<paragraph pageId="0" pageNumber="997">
<emphasis bold="true" pageId="0" pageNumber="997">Figure 6.</emphasis>
Type localities (stars) and other collection sites (circles) of
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
Group species.
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
Map of New Guinea and its satellite islands, showing the positions of the inset maps and the type locality of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mintoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mintoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. mintoni</emphasis>
</taxonomicName>
(magenta), an endemic of Sudest Island, Milne Bay Province, PNG. Scale = 500 km.
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
Papua New Guinea and adjacent areas of West New Guinea, Indonesia, with white lines demarcating provincial boundaries. Scale = 250 km.
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
West Papua Province, Indonesia (WP), showing the type locality of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loennbergii" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loennbergii">
<emphasis italics="true" pageId="0" pageNumber="997">T. loennbergii</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(orange). Other species treated in this report are
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
(dark blue),
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. lamingtoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="lamingtoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. lamingtoni</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(yellow),
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. spilorhynchus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="spilorhynchus">
<emphasis italics="true" pageId="0" pageNumber="997">T. spilorhynchus</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
(red),
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nymani" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nymani">
<emphasis italics="true" pageId="0" pageNumber="997">T. nymani</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">comb. nov.</emphasis>
(neon green),
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. vertebralis" order="Squamata" pageId="0" pageNumber="997" rank="species" species="vertebralis">
<emphasis italics="true" pageId="0" pageNumber="997">T. vertebralis</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
(pink), and
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. atratus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="atratus">
<emphasis italics="true" pageId="0" pageNumber="997">T. atratus</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="997">sp. nov.</emphasis>
(light blue). Additional related species in the
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
clade are
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. goodenoughensis" order="Squamata" pageId="0" pageNumber="997" rank="species" species="goodenoughensis">
<emphasis italics="true" pageId="0" pageNumber="997">T. goodenoughensis</emphasis>
</taxonomicName>
(brown),
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. mattisoni" order="Squamata" pageId="0" pageNumber="997" rank="species" species="mattisoni">
<emphasis italics="true" pageId="0" pageNumber="997">T. mattisoni</emphasis>
</taxonomicName>
(violet),
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. nigrescens" order="Squamata" pageId="0" pageNumber="997" rank="species" species="nigrescens">
<emphasis italics="true" pageId="0" pageNumber="997">T. nigrescens</emphasis>
</taxonomicName>
(black),
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. pachysomus" order="Squamata" pageId="0" pageNumber="997" rank="species" species="pachysomus">
<emphasis italics="true" pageId="0" pageNumber="997">T. pachysomus</emphasis>
</taxonomicName>
(dark green), and
<taxonomicName family="Elapidae" genus="T." kingdom="Animalia" lsidName="T." order="Squamata" pageId="0" pageNumber="997" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="997">T.</emphasis>
</taxonomicName>
&quot;
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
&quot; Clade 4 (white). The type locality of the recently described dubious taxon
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. longhagen" order="Squamata" pageId="0" pageNumber="997" rank="species" species="longhagen">
<emphasis italics="true" pageId="0" pageNumber="997">T. longhagen</emphasis>
</taxonomicName>
(tan) is also indicated. Scale = 250 km. Provinces of PNG are Central (Ce), Chimbu (Ch; also spelled Simbu), Gulf (Gu), Hela (He; created in 2012), Eastern Highlands (EH), East Sepik (ES), Enga (En), Jiwaka (Ji; created in 2012), Madang (Ma), Milne Bay (MB), Morobe (Mo), National Capital District (NCD), Oro (Or; also known as Northern Province); Sandaun (Sa; formerly West Sepik), Southern Highlands (SH), Western (We), and Western Highlands (WH). Provinces of West New Guinea are Highland Papua (HP; created in 2022), Papua (Pa), and South Papua (SP; created in 2022). Sites of sympatry are Wau (W; neon green + pink) with two species and Garaina (G; neon green + red + yellow) with three species.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="ecological notes">
<paragraph pageId="0" pageNumber="997">Ecological notes.</paragraph>
<paragraph pageId="0" pageNumber="997">
Three individuals from Mt. Obree (elevation ca. 880 m) were obtained from local villagers, who presumably discovered the snakes while gardening. One specimen came from near a stream in primary rainforest (1570 m), and one more was found on the forest floor in primary rainforest on a ridge top (1680 m). At the time of collection, the understory on this ridge was not dense and contained a thick duff layer with a tight root network at the surface; the canopy was 25-30 m above the forest floor, with large emergent araucarians (
<taxonomicName class="Pinopsida" family="Araucariaceae" genus="Araucaria" kingdom="Plantae" lsidName="Araucaria" order="Pinales" pageId="0" pageNumber="997" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="997">Araucaria</emphasis>
</taxonomicName>
sp.,
<taxonomicName class="Pinopsida" family="Araucariaceae" kingdom="Plantae" lsidName="" order="Pinales" pageId="0" pageNumber="997" phylum="Tracheophyta" rank="family">Araucariaceae</taxonomicName>
) reaching heights of 50 m. Farther downslope, near the stream, the forest had a much denser understory. These specimens were found active on the forest floor at 12:00 h and in mid-afternoon (~16:00 h), respectively.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="remarks">
<paragraph pageId="0" pageNumber="997">Remarks.</paragraph>
<paragraph pageId="0" pageNumber="997">
We retain
<taxonomicName authorityName="Kraus &amp; Kaiser &amp; O’Shea" authorityYear="2022" baseAuthorityName="Kraus &amp; Kaiser &amp; O’Shea" baseAuthorityYear="2022" class="Reptilia" family="Elapidae" genus="Apistocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apistocalamus pratti" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="pratti">
<emphasis italics="true" pageId="0" pageNumber="997">Apistocalamus pratti</emphasis>
</taxonomicName>
(BMNH 1946.1.17.53, Fig.
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Head of holotype of Apistocalamus pratti (BMNH 1946.1.17.53) from Dinawa, Central Province, PNG, in both photographic and line-drawing illustrations. A, A ' dorsal, B, B ' right lateral, C, C ' left lateral, and D, D ' ventral views." figureDoi="10.3897/vz.72.e89647.figure7" httpUri="https://binary.pensoft.net/fig/768173" pageId="0" pageNumber="997">7</figureCitation>
) in the synonymy of
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
because the sole specimen shares with the holotype of the latter the two diagnostic features of having two intergenials separating the posterior genials and a yellow venter with a mid-ventral row of brown spots. It differs from the holotype of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
in being smaller (330 vs. 490 mm SVL), having slightly fewer ventrals (186 vs. 189) and subcaudals (40 vs. 46), having a single postocular (vs. 2), and having a small, indistinctly defined yellow spot on each prefrontal. The last feature merely reflects variation seen in some of the smaller specimens of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae." order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae.">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae.</emphasis>
</taxonomicName>
The other differences are trivial and not diagnostic.
</paragraph>
<caption doi="10.3897/vz.72.e89647.figure7" httpUri="https://binary.pensoft.net/fig/768173" pageId="0" pageNumber="997" start="Figure 7" startId="F7">
<paragraph pageId="0" pageNumber="997">
<emphasis bold="true" pageId="0" pageNumber="997">Figure 7.</emphasis>
Head of holotype of
<taxonomicName authorityName="Kraus &amp; Kaiser &amp; O’Shea" authorityYear="2022" baseAuthorityName="Kraus &amp; Kaiser &amp; O’Shea" baseAuthorityYear="2022" class="Reptilia" family="Elapidae" genus="Apistocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apistocalamus pratti" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="pratti">
<emphasis italics="true" pageId="0" pageNumber="997">Apistocalamus pratti</emphasis>
</taxonomicName>
(BMNH 1946.1.17.53) from Dinawa, Central Province, PNG, in both photographic and line-drawing illustrations.
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">A</emphasis>
' dorsal,
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">B</emphasis>
' right lateral,
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">C</emphasis>
' left lateral, and
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="997">D</emphasis>
' ventral views.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="997">
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus loriae" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus loriae</emphasis>
</taxonomicName>
is unique among members of the
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
Group in having the posterior genials completely separated from contact with each other by two intervening intergenials aligned in the midline of the chin. This character state is among the more distinctive features that diagnose any of the
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
Group species. Furthermore, this feature occurs in only a small, contiguous portion of the entire geographical range occupied by the group. In our view, it provides compelling evidence for the unity of this species. However, we note that the sample size for this species is small (
<emphasis italics="true" pageId="0" pageNumber="997">n</emphasis>
= 19) and almost one-third of those specimens come from a single area (on the NW slope of Mt. Obree); all other localities are represented by only 1-3 specimens. Consequently, it is difficult to assess the importance of some of the morphological and geographic variation seen in this sample. We here consider four points in question that deserve further assessment once sufficiently large specimen series become available.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="ventral colour pattern">
<paragraph pageId="0" pageNumber="997">Ventral colour pattern.</paragraph>
<paragraph pageId="0" pageNumber="997">
This character is variable. Three specimens (AMNH R-59067, BMNH 1935.5.10.174, PNGM 23158) from Mafulu and from near Fane (9 km apart) show an expansion of brown pigment on each ventral that is not seen in any other specimens. In particular, the anterior edge of each ventral is adorned with a narrow bar of brown. Other specimens are either uniformly yellow ventrally (37%) or yellow with a row or scattering of brown dots mid-ventrally (42%); one specimen (NMW 27383.2) is uniformly pale brown ventrally, though we suspect this may be an artifact of preservation or storage. Of the three specimens from Mafulu and Fane, two still retain the central row of brown dots but the brown colouration extends laterally to these dots. Assuming these specimens are true
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
, this extends the range of colour-pattern variation seen in the species. We believe these are true
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
, both because they have the characteristic medial separation of the posterior genials by two intergenials as well as the fact that the medial brown dots are still evident in two of these three specimens. Furthermore, the sample of six specimens from Mt. Obree also makes clear that the degree of brown markings on the venter is variable within this species: one of these specimens has a yellow venter with a complete row of mid-ventral brown spots, two have yellow venters with some mid-ventral brown spotting, and three have uniformly yellow venters. Variation in degree of brown ventral spotting does not appear to be ontogenetic since the largest adults can be either uniformly yellow or have a complete series of brown spots.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="ventral counts">
<paragraph pageId="0" pageNumber="997">Ventral counts.</paragraph>
<paragraph pageId="0" pageNumber="997">
Male ventral counts in our sample cover an unusually wide range (162-197) and might be thought to possibly reflect the presence of more than one species in our sample. However, these values seem to vary geographically, with two of the three highest counts (193, 194, 197) seen in the two northernmost samples (Fane, Mafulu) and the two lowest counts in the southernmost samples from Agaun, Milne Bay Province (162) and nearby Mt. Obree (169). Furthermore, the six males from Mt. Obree, Central Province, themselves show a range of variation (169-189) that encompasses 55% of the total variation seen in the entire sample of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
, suggesting that considerable variation in male ventral counts does indeed characterize this species and may not in itself be evidence of multiple species. This interpretation could possibly change when larger sample sizes become available, but the overlap in counts seen in the Agaun and Mt. Obree samples, coupled with the broad variation seen in the latter, makes this seem unlikely.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="geography">
<paragraph pageId="0" pageNumber="997">Geography.</paragraph>
<paragraph pageId="0" pageNumber="997">
The large majority of specimens was collected on the southern versant of the central Owen Stanley Range. However, one specimen (BPBM 44892) came from the northern versant of these mountains in Oro Province. This is an unexpected locality for what appears to be a southern-versant species, for two reasons. Firstly, two other species of the
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
Group also occur in this general area of Oro Province (see below), and one of those is restricted to the northern versant. However, current evidence suggests that these species are segregated elevationally in this area, with the sole
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
specimen having been collected at an elevation of 1260 m, the second species occurring in hill forest from 100-940 m, and the third appearing to be a high-elevation form found at 1660-1850 m. Secondly, even if these species all segregate along elevational gradients, it remains uncertain how
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
would have crossed the formidable barrier presented by the Owen Stanley Range (elevation&gt; 2000 m in this region). It could possibly have spread around these mountains from the south, through the low-elevation Musa Divide. If that is true, then additional surveys of this poorly explored region should reveal a more extensive presence of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
at middle elevations of the Owen Stanley Range in Oro Province.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="997" type="provenance">
<paragraph pageId="0" pageNumber="997">Provenance.</paragraph>
<paragraph pageId="0" pageNumber="997">
Two specimens (NMW 27383.1-2) are of ambiguous provenance. These were sent to Vienna by the anthropologist and explorer Rudolf
<normalizedToken originalValue="Pöch">Poech</normalizedToken>
(1870-1921), who worked in German New Guinea from 1904-06 (
<bibRefCitation DOI="https://doi.org/10.2307/­17­76812" author="Poech, R" journalOrPublisher="The Geographical Journal" pageId="0" pageNumber="997" pagination="609 - 616" refId="B37" refString="Poech, R, 1907. Travels in German, British, and Dutch New Guinea. The Geographical Journal 30 (6): 609 - 616, DOI: https://doi.org/10.2307/-17-76812" title="Travels in German, British, and Dutch New Guinea." url="https://doi.org/10.2307/­17­76812" volume="30" year="1907">
<normalizedToken originalValue="Pöch">Poech</normalizedToken>
1907
</bibRefCitation>
). The locality for these specimens (&quot;Astrolabe Gebirge&quot;) can be interpreted as referring to the mountains visible from Astrolabe Bay (potentially either the Finisterre or Adelbert Mts.) in Madang Province, or to the Astrolabe Range south of the Owen Stanley Range in Central Province. The former interpretation is consistent with the fact that most of
<normalizedToken originalValue="Pöch’s">Poech's</normalizedToken>
work was done in German New Guinea, including the area now contained within Madang Province. Against this is the fact that all
<taxonomicName authorityName="Boulenger" authorityYear="1896" class="Reptilia" family="Elapidae" genus="Toxicocalamus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Toxicocalamus" order="Squamata" pageId="0" pageNumber="997" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="997">Toxicocalamus</emphasis>
</taxonomicName>
in the
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
Group otherwise collected from this region are not
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
and do not match the NMW specimens. Contrarily, if the locality is interpreted to refer to the Astrolabe Range in Central Province, this would make geographical and morphological sense, because these specimens would then be from within 25 km of the type locality of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
and are morphologically unremarkable
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
specimens.
<normalizedToken originalValue="Pöch">Poech</normalizedToken>
is known to have visited nearby Port Moresby for a few days before his departure from New Guinea, and it is possible that the specimens were on hand and given to him there by his hosts. In any event, these two specimens are morphologically and geographically consistent with their provenance being from the Astrolabe Range, Central Province. If they truly did come from somewhere in Madang Province, then they are morphologically unique for that region and would suggest that either
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
has a much wider range than currently evident (&gt; 350 km from the nearest known locality for the species) or that a morphologically similar species occurs in the Madang region that has not been collected since
<normalizedToken originalValue="Pöch’s">Poech's</normalizedToken>
time. Some specimens were indeed sent to Vienna from Port Moresby in the time frame under consideration, but there is no further information in the records of the Vienna Museum to determine what the shipment may have included. We tentatively adopt the position that these specimens came from Central Province, and we have included them in the diagnosis of
<taxonomicName family="Elapidae" kingdom="Animalia" lsidName="T. loriae" order="Squamata" pageId="0" pageNumber="997" rank="species" species="loriae">
<emphasis italics="true" pageId="0" pageNumber="997">T. loriae</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>